Coprophanaeus ensifer

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Coprophanaeus ensifer
C. ensifer.jpg
Coprophanaeus ensifer in Pernambuco, Brazil
Scientific classification
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Species:
C. ensifer
Binomial name
Coprophanaeus ensifer
(Germar, 1821)
Synonyms
  • Copris ensifer Germar, 1821
  • Megaphanaeus ensifer Germar, 1821
  • Phanaeus ajax Sturm, 1826
  • Phanaeus ducalis Castelnau, 1840

[1]

Coprophanaeus ensifer is a large South American species of beetle belonging to the family Scarabaeidae. [2] This species is necrophagous and builds burrows near or on animal carcasses to dismember the flesh of decaying bodies and bring it to its burrow to feed. Both females and males help build the burrow and feed. It is characterized by its iridescent colors and a horn that is similar in shape and size in females and males. It uses its horn to tear apart carcasses and to fight with other individuals, with male-male fighting occurring more often. However, females also fight to determine a variety of characteristics of the opposing male. This species may be of importance in forensic science due to its destructive behavior on decaying bodies, especially in areas of Brazil where homicide rates are high.

Phylogeny

C. ensifer vary in color and distribution, which has created uncertainty and barriers in its taxonomy within the Coprophanaeus genus. Based on its mitogenome, the species was placed in the Phaneini tribe, which forms a clade of Scarabaeinae with other tribes like the Coprini tribe. Among species in this clade, most reside in South America except for Sarophorus sp., which is found in South Africa. Based on this, researchers have suggested Scarabaeinae beetles came to South America from Africa through colonization of the common ancestor of Sarophorus sp. and species in the clade with C. ensifer. [3]

Distribution and habitat

This species is found widely in eastern, central and southern Brazil, far northeastern Argentina, eastern Bolivia and Paraguay, occurring up to about 780 m (2,560 ft) above sea level. [2] [4] [5] [6] Most of its range is in the Caatinga, Cerrado and similar habitats, but in some regions its range also extends into the Atlantic forest and the Amazon rainforest. [2] [5]

A relates species, C. lancifer occupies northern Brazil, limiting the distribution of C. ensifer in those regions. [7] C. lancifer was also thought to occupy the Juína region, but upon genetic investigation, it was found that individuals in that region are more closely related to C. ensifer, suggesting that the distribution of C. ensifer is wider than previously thought. [8]

A study found that in Northeast Brazil, C. ensifer is more active during the rainy season, when temperatures drop. This preference can be explained by their large body. At high temperatures during dry seasons, it could be difficult to maintain a proper water balance because a large amount of water can be evaporated from their bodies. This could also explain why these beetles are crepuscular, or most active at dawn and dusk. [9]

Building burrows

In many cases, both females and males work together to build a burrow. Pairs that build burrows together are usually larger than other conspecifics. Females mainly built the burrow by pushing soil while males bring the soil up to the surface. The burrow is usually built near a carcass so that females and males can break it down to bring it inside the burrow by rolling the pieces of food. [7] Cooperation between males and females suggests that parental care is offered by both sexes. [10]

In other cases, females and males build burrows individually despite having mated with other individuals. Occasionally, small individuals build burrows next to burrows built by a pair. [7]

Although the effects of bioturbation, or moving and mixing soil, caused by building burrows has not been empirically studied, it has been suggested that this could have an effect on the soil itself by increasing aeration and the ability for water to penetrate. [11]

Diet

Compared to other Scarabaeidae species who are coprophagous, this species is necrophagous, which means it feeds on carrion, or the decaying body of dead animals. It has even been reported to bury a dead snake to feed on it. [12] A carcass goes through various stages of decomposition and C. ensifer is most attracted to the body when it is actively decomposing in the initial stages of decomposition. [10]

By breaking down carrion and moving it or burying it, nutrient cycling may occur where nutrients that were part of the decaying carcass reenter the soil, affecting plant growth. [11]

It has been suggested that the difference in diet in C. ensifer compared to other species in the Scarabaeidae family evolved in the Pleistocene because of the extinction of large mammals, so the beetles were forced to change their diet to decrease competition and ensure survival. [9]

These beetles find decomposing carcasses through the detection of volatiles released during the decomposition process. A study found that females always arrived first to a pig carcass after the sun set, while males arrived later. Once the beetles arrive at the carcass, males bury themselves while females feed from it using their horn. The difference in arrival time suggest that females first find a suitable resource to feed and for oviposition while males arrive later to mate with females. Another potential explanation could be that males take advantage of females who have started building burrows, so that they can avoid digging tunnels. [10]

Fighting behavior

Both females and males engage in fighting activity using their horns. There can be male-male, female-female, or male-female fighting in these beetles. In burrows, individuals use their horn to carry and drop the other but on the surface of burrows, individuals fight by pushing each other. Males are more aggressive than females. [7]

Female fighting against males may simply occur to refuse copulation after they have already copulated. However, another suggestion is that fighting can be used as a way to determine mates. If horns act as an ornament to attract females, fighting could be a way for females to gauge which males to mate with. If females can also decide which sperm to use to fertilize their eggs, engaging in fights after copulation could also serve to detect which sperm to use. When males fight, the larger male always wins, so this may also be a mechanism for mate choice. Lastly, males may also determine which female to mate with based on fighting. Larger females may be more beneficial to mate with because they can protect burrows, so fighting can be a way for males to determine female size. [7]

Genetics

The size of the mitogenome of C. ensifer is 14,964 bp organized into rRNA, tRNA, and protein coding genes, which is similar to species in the Coleptera order. [3] C. ensifer, along most of Scarabaeinae species, are diploid (2n=20). This species specifically, has a XY sex-determining mechanism where the X and Y chromosomes do not exhibit any different configurations observed in other species. [13]

Copulation behavior

Four stages of copulation have been defined for Neotropical Scarabaeinae beetles. The first stage involves the male approaching the female, which is usually around a carcass or burrow. In the second stage, the male detects the female. The third stage involves the male mounting on the female. [14] It has been observed that while the mounting occurs, the male strokes the abdominal tergum with his legs so that the female can reveal her genital opening. [7] During the last stage, the males insert their aedeagus in the female genital opening. [14]

While all males attempt to mate with females, larger males are more successful and among males, copulations are more successful when attempting with a smaller female. [7] Copulation activity mainly takes place underground in burrows and tunnels and lasts around one to two minutes. [14]

Description

Adults typically are 30–56 mm (1.2–2.2 in) long, [2] occasionally even reaching 65 mm (2.6 in). [15] This makes C. ensifer and the closely related Amazonian C. lancifer some of the largest dung beetles in the world (together with certain Catharsius and Heliocopris ) and the largest in the Americas. [2] [4] [5] This beetle has a horn on the head that is of similar size in males and females. It is usually dark metallic green, often with some blue reflections, but it can occasionally be all metallic blue. [4] [5] [16]

Sexual dimorphism

Female and male C. ensifer do not exhibit pronounced sexual dimorphism. The main differences are that females do not have a spiniform process in the pronotal concavity compared to males, but they have a protarsi that males do not have. [2]

Horn

C. ensifer has a long horn with a pair of processes projecting up and two other pairs of processes projecting forward. Although the body size of females and males is similar, the male horns are relatively larger than female horns. This could be because males have to compete for females.  The horn has been suggested to play roles in sexual selection, defense, as a tool for building burrows, or during conflict between individuals. Specifically, females may have evolved the horn to defend their burrow since they can only have a limited number of offspring due to their small reproductive system. [7]

Chromatic variations

C. ensifer vary in color depending on their location. Those near the Amazon Rainforest exhibit bluer and darker colors while those near in the Atlantic Forest have green and brighter patterns. In the Cerrado, or the Brazilian savanna, beetles exhibit a variety of reflexes ranging from green to blue to red. Although the significance of the iridescence of C. ensifer has not been examined, it has been suggested to be a mechanism for a variety of things like defense against predators, sexual selection, or signaling. [5]

Flight

Coprophanaeus beetles are known to fly for a short time after sunset. The short period of flight could be due to their diet. Carcasses are unpredictable, so it might be more efficient for the beetles to fly for a short amount of time to determine the presence of a carcass instead of using energy to fly long periods of time continuously looking for one. [17]

Relevance in Forensics

Because of their necrophagous diet, this species has been regarded as forensically important. Homicide rates in Brazil have increased over the years, and necrophagous species have been associated with determining post-mortem information about cadavers. Although the distribution and diet of these species is understood, the actual use of the information in investigations is not well established in Brazil’s forensic systems. [18]

Related Research Articles

<span class="mw-page-title-main">Scarabaeidae</span> Family of beetles

The family Scarabaeidae, as currently defined, consists of over 35,000 species of beetles worldwide; they are often called scarabs or scarab beetles. The classification of this family has undergone significant change in recent years. Several groups formerly treated as subfamilies have been elevated to family rank, and some reduced to lower ranks. The subfamilies listed in this article are in accordance with those in Catalog of Life (2023).

<span class="mw-page-title-main">Dung beetle</span> Informal group of insects

Dung beetles are beetles that feed on feces. Some species of dung beetles can bury dung 250 times their own mass in one night.

<span class="mw-page-title-main">Hercules beetle</span> Species of beetle

The Hercules beetle is a species of rhinoceros beetle native to the rainforests of southern Mexico, Central America, South America, and the Lesser Antilles. It is the longest extant species of beetle in the world, and is also one of the largest flying insects in the world.

<i>Phanaeus vindex</i> Species of beetle

Phanaeus vindex, also known as a rainbow scarab, is a North American species of true dung beetle in the family Scarabaeidae. It is found in eastern and central United States and northern Mexico. It is the most widespread species of Phanaeus in the United States and it has a wide habitat tolerance. It may hybridize with the generally less common P. difformis.

<span class="mw-page-title-main">Scarabaeinae</span> Subfamily of beetles

The scarab beetle subfamily Scarabaeinae consists of species collectively called true dung beetles. Most of the beetles of this subfamily feed exclusively on dung. However, some may feed on decomposing matter including carrion, decaying fruits and fungi. Dung beetles can be placed into three structural guilds based on their method of dung processing namely rollers (telecoprids), dwellers (endocoprids) and tunnelers (paracoprids). Dung removal and burial by dung beetles result in ecological benefits such as soil aeration and fertilization; improved nutrient cycling and uptake by plants, increase in Pasture quality, biological control of pest flies and intestinal parasites and secondary seed dispersal. Well-known members include the genera Scarabaeus and Sisyphus, and Phanaeus vindex.

<i>Nicrophorus quadripunctatus</i> Species of beetle

Nicrophorus quadripunctatus is a species of burying beetle that predominates in East Asia. First described by German entomologist Ernst Kraatz in 1877, this beetle has since been the subject of much scientific inquiry—particularly concerning its parental care. Like other burying beetles, N. quadripunctatus inhabit small, vertebrate animal carcasses. This environment provides the beetles with the requisite nutrients for themselves and their offspring. To limit resource theft and predation, the carcass is buried underground. For additional protection, a single, dominant male-female pair guards the carcass cooperatively.

<i>Catharsius</i> Genus of beetles

Catharsius is a genus of dung beetles in the tribe Coprini in the scarab family.

<i>Coprophanaeus</i> Genus of beetles

Coprophanaeus is a genus in the family Scarabaeidae. The genus is almost entirely Neotropical, with a single species, C. pluto, ranging into southernmost Texas in the United States. They are medium-sized to large beetles, with the South American C. ensifer and C. lancifer sometimes exceeding 5 cm (2 in) in length, making these two some of the largest dung beetles in the world and the largest in the Americas. They often have a horn on the head, and are typically a bright metallic color, most often blue or green, or black. These diurnal or crepuscular beetles are excellent diggers and good fliers.

<i>Heliocopris</i> Genus of beetles

Heliocopris is a genus of Scarabaeidae or scarab beetles in the superfamily Scarabaeoidea. Forty-seven of the fifty-two known species are found in Africa, but a few are found in southern and southeast Asia.

<i>Bolitotherus</i> Species of beetle

Bolitotherus cornutus is a North American species of darkling beetle known as the horned fungus beetle or forked fungus beetle. All of its life stages are associated with the fruiting bodies of a wood-decaying shelf fungus, commonly Ganoderma applanatum, Ganoderma tsugae, and Ganoderma lucidum.

<i>Onthophagus taurus</i> Species of beetle

Onthophagus taurus, the taurus scarab, is a species of dung beetle in the genus Onthophagus and the family Scarabaeidae. Also known as the bull-headed dung beetle, it is a species that specializes in cattle dung and is widely utilized to maintain clean pastures, making it agriculturally valuable. These beetles are typically 8–10 millimetres (0.31–0.39 in) in size. The males of this species exhibit distinct characteristics: large “major” males possess long, sweeping, curved horns resembling those of a longhorn bull, while small “minor” males have tiny horns that project upward from the back of their heads. Females, on the other hand, lack horns. These small beetles are oval shaped, the color is usually black or reddish brown. Sometimes the pronotum has a weak metallic sheen.

<i>Phanaeus</i> (beetle) Genus of beetles

Phanaeus, the rainbow scarabs, is a genus of true dung beetles in the family Scarabaeidae, ranging from the United States to northern Argentina, with the highest species richness in Mexico. Depending on species, they can inhabit a wide range of habitats, from tropical to temperate climates and deserts to rainforests. In those living in relatively arid places adults are primarily active during the wet season and those living in relatively cold places are primarily active during the summer. They are excellent diggers and good fliers.

<i>Sulcophanaeus imperator</i> Species of beetle

Sulcophanaeus imperator is a brightly colored species of dung beetle belonging to the family Scarabaeidae. This diurnal, coprophagous beetle is native to south-central South America and generally common. It is paracoprid, meaning that adults dig tunnels into the soil under the food source and move parts of the food source to a nest chamber where the eggs are laid.

<span class="mw-page-title-main">Necrophage</span> Organism that consumes dead animal matter

Necrophages are organisms that obtain nutrients by consuming decomposing dead animal biomass, such as the muscle and soft tissue of carcasses and corpses. The term derives from Greek nekros, meaning 'dead', and phagein, meaning 'to eat.' Mainly, necrophages are species within the phylum Arthropoda; however, other animals, such as gastropods and Accipitrimorphae birds have been noted to engage in necrophagy.

<i>Oxysternon festivum</i> Species of beetle

Oxysternon festivum is a species of dung beetle of the scarab beetle family. It is a common, diurnal species from northeastern South America and Trinidad where it mostly inhabits forest.

<span class="mw-page-title-main">Coprini</span> Tribe of beetles

Coprini is a tribe of scarab beetles, in the dung beetle subfamily (Scarabaeinae). Scholtz et al. describe them as tunnellers that are shiny black, of moderate to large size and with a strongly convex shape. They also, however state that the grouping based on these characteristics has little phylogenetic validity, and the placement of several genera in this and related tribes is likely to change.

<span class="mw-page-title-main">Phanaeini</span> Tribe of beetles

Phanaeini is a tribe of dung beetles in the family Scarabaeidae. There are about 12 genera and 200 described species in Phanaeini. They are native to the Americas with the highest species richness in the Neotropics. They are mostly coprophagous or necrophagous, but some of the least known genera appear to be myrmecophilous. They are medium-sized to large beetles, often with bright metallic colors, and often with horns on their heads.

<i>Euoniticellus intermedius</i> Species of beetle

Euoniticellus intermedius is a species of dung beetle in the family Scarabaeidae. E. intermedius is native to Southeastern Africa but has spread to the United States, Mexico, and Australia. E. intermedius acts as an important agricultural agent due to its improvement of soil quality and removal of parasitic pests.

<i>Oxysternon conspicillatum</i> Species of beetle

Oxysternon conspicillatum is a species of dung beetle in the family Scarabaeidae. It is found in both evergreen and semi-deciduous mesic forests, including disturbed habitats, ranging from near sea level to an altitude of 3,000 m (9,800 ft) in the Amazon basin and Chocó of tropical South America and Panama, possibly extending as far west as the border region with Costa Rica. Both adults and young of this common and widespread beetle primarily feed on dung, but the species has also been recorded feeding on dead animals.

<i>Coprophanaeus lancifer</i> Species of beetle

Coprophanaeus lancifer is a large species of beetle belonging to the family Scarabaeidae.

References

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